A Reappraisal of Phylogentic Relationships in the Monocotyledon Family Hydrocharitaceae (Alismatidae)

Abstract

The diverse, aquatic Hydrocharitaceae have defied stable classification for nearly two centuries. Anatomical and morphological convergence characterize many aquatic plants and undoubtedly have hindered the ability of researchers to ascertain accurately those features representing reliable phylogenetic markers within Hydrocharitaceae. Most prior classifications of the family have emphasized few characters to define major taxonomic subdivisions (i.e., they were highly artificial). Previous studies using molecular data have shown that DNA sequences provide novel indications of phylogeny not indicated previously by morphologically based classifications; however, they have not yet recommended alterations to the classification for the family. We conducted a more comprehensive phylogenetic study of Hydrocharitaceae to better elucidate evolutionary relationships among the genera that in turn could be used to provide insight for improvements in classification. We analyzed different data sets (55 morphological characters; chloroplast rbcL, matK, trnK intron sequences; nuclear ribosomal ITS region sequences) singly and in various combinations using maximum parsimony and maximum likelihood methods of phylogenetic reconstruction. Phylogenetic analysis of combined data yielded a fully resolved tree depicting four well-supported, major clades within Hydrocharitaceae. We use these results to propose a phylogenetic classification of Hydrocharitaceae recognizing four subfamilies that correspond to these clades: Anacharidoideae, Hydrilloideae, Hydrocharitoideae, and Stratioideae. Phylogenetic analysis also indicated the pattern of derivation with respect to submersed lifeforms, hydrophilous pollination, and marine habitation in the family. Character reconstructions indicated that several features, (e.g., ovule type; occurrence of detaching male flowers), once thought to provide strong phylogenetic markers in Hydrocharitaceae, actually are highly homoplasious and have acutely mislead past attempts at classification of the family.